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Universidad Nacional Del Comahue Centro Regional Universitario Bariloche Universidad Nacional del Comahue Centro Regional Universitario Bariloche Título de la Tesis Microanatomía y osteohistología del caparazón de los Testudinata del Mesozoico y Cenozoico de Argentina: Aspectos sistemáticos y paleoecológicos implicados Trabajo de Tesis para optar al Título de Doctor en Biología Tesista: Lic. en Ciencias Biológicas Juan Marcos Jannello Director: Dr. Ignacio A. Cerda Co-director: Dr. Marcelo S. de la Fuente 2018 Tesis Doctoral UNCo J. Marcos Jannello 2018 Resumen Las inusuales estructuras óseas observadas entre los vertebrados, como el cuello largo de la jirafa o el cráneo en forma de T del tiburón martillo, han interesado a los científicos desde hace mucho tiempo. Uno de estos casos es el clado Testudinata el cual representa uno de los grupos más fascinantes y enigmáticos conocidos entre de los amniotas. Su inconfundible plan corporal, que ha persistido desde el Triásico tardío hasta la actualidad, se caracteriza por la presencia del caparazón, el cual encierra a las cinturas, tanto pectoral como pélvica, dentro de la caja torácica desarrollada. Esta estructura les ha permitido a las tortugas adaptarse con éxito a diversos ambientes (por ejemplo, terrestres, acuáticos continentales, marinos costeros e incluso marinos pelágicos). Su capacidad para habitar diferentes nichos ecológicos, su importante diversidad taxonómica y su plan corporal particular hacen de los Testudinata un modelo de estudio muy atrayente dentro de los vertebrados. Una disciplina que ha demostrado ser una herramienta muy importante para abordar varios temas relacionados al caparazón de las tortugas, es la paleohistología. Esta disciplina se ha involucrado en temas diversos tales como el origen del caparazón, el origen del desarrollo y mantenimiento de la ornamentación, la paleoecología y la sistemática. Además, dada la relevancia de la microestructura ósea, las descripciones histológicas también se han incluido en recientes estudios anatómicos y sistemáticos en tortugas fósiles. En este estudio, para abordar problemas concernientes a varios aspectos de la paleobiología de las tortugas, fueron examinamos alrededor de 400 cortes delgados pertenecientes a unas 200 placas que corresponden a más de 30 especies fósiles y actuales. Para llevar a cabo el estudio de estos cortes delgados, se propusieron los siguientes objetivos: caracterizar la microanatomía y la osteohistología de la armadura dérmica de distintas especies de tortugas del Jurásico, Cretácico y Cenozoico de Argentina, así también como de Testudinata actuales (Pleurodira y Criptodira); identificar variaciones microanatómicas y osteohistológicas a nivel inter-elementos dentro de la armadura dérmica; reconocer posibles variaciones microanatómicas y osteohistológicas entre distintos linajes de Testudinata y establecer el valor taxonómico de dichos caracteres; determinar el tipo y patrón de histogénesis del caparazón de los Testudinata; evaluar la posible correlación entre la organización interna del caparazón y la adaptación a distintos tipos de hábitats (marino, dulceacuícola y terrestre); e identificar variaciones ontogenéticas en el desarrollo del caparazón de los Testudinata. Por último, se propone comparar los 2 Tesis Doctoral UNCo J. Marcos Jannello 2018 resultados obtenidos con los estudios publicados enfocados en la histología ósea de los Testudinata. Las principales contribuciones de esta tesis a la paleohistología del caparazón de las tortugas se exponen a continuación. En primer lugar, se registraron nuevos patrones de variación histológica intra- e inter-elemento y se corroboraron los patrones conocidos previamente. Se encontró una nueva e importante variación de la histología ósea entre las placas periféricas y el resto de las placas del caparazón. En segundo lugar, con respecto a la histogénesis, el aporte de osificación metaplásica no solo se observa en forma variable entre los distintos taxones y los distintos elementos del caparazón, sino también entre distintas regiones dentro de los elementos. La osificación metaplásica ocurre únicamente en la corteza externa, salvo en contadas excepciones donde también está presente en la corteza interna. Durante la ontogenia, la osificación metaplásica ocurre temprano y nunca surge en etapas ontogenéticas tardías. Los diferentes patrones reportados en el desarrollo y la variación en cuanto al aporte de la osificación metaplásica de los Testudinata no muestran una clara señal que agrupe a taxones. Si bien es conocido que el crecimiento en espesor de las placas durante la ontogenia ocurre hacia la región externa, se reporta por primera vez, sólo en algunas especies, un nuevo patrón de crecimiento, el cual es inverso al anterior (i.e. crecimiento hacia la región interna). Se describen y corroboran los dos mecanismos básicos para la formación y el mantenimiento de la ornamentación externa entre los Testudinata; por un lado, depositación con variación en las tasas locales de formación, por el otro lado reabsorción ósea y nueva depositación de tejido óseo. No se encontró una relación evidente, en principio, entre el tipo o dimensión de la ornamentación y el patrón de formación y mantenimiento de la misma. En tercer lugar, contrario a lo establecido en varias contribuciones previas, se sugiere que no existe una correlación entre el hábitat y la histología del caparazón en los Testudinata. Finalmente, la microanatomía y la osteohistología del caparazón de las tortugas son fuente de información para estudios sistemáticos. Los caracteres histológicos se distribuyen en categorías taxonómicas menos inclusivas (i.e., especies, géneros y, eventualmente, familias). Se reconocen caracteres singulares en la corteza interna del caparazón de todas las especies estudiadas del género Podocnemis. Se reconocieron rasgos con potencial utilización en sistemática en Prochelidella cerrobarcinae, Trapalcochelys sulcata, Mendozachelys wichmanni y Pelomedusoides gen. et sp. indeterminado. 3 Tesis Doctoral UNCo J. Marcos Jannello 2018 Summary Unusual bone structures between vertebrates, like the giraffe´s long neck or the T- shaped skull of the hammerhead shark, have interested scientists since a long time ago. Testudinata clade represent one of the most fascinating and enigmatic group of amniotes. Their peculiar body plan, characterized by the development of a bony shell that encloses the pectoral girdles within the rib cage, has actually persisted since the Late Triassic and allowed them to adapt successfully to different environments (e.g., terrestrial, freshwater, coastal and even pelagic marine). Their capability in facilitating different ecological niches, important taxonomical diversity and particular body plan make Testudinata a highly interesting model of studies in vertebrates. Paleohistological studies of turtle bones in extinct taxa have proven to be a very important tool for addressing several issues, including: shell origin, development origin and maintenance of shell ornamentation, paleoecology and systematics. Moreover, given the relevance of the bone microstructure, histological descriptions have also been included in recent anatomical and systematic studies of extinct turtles. In this survey, 400 slides from more than 200 elements belonging to more than 30 species of extinct and extant turtles were examined to address several issues about their paleobiology.The aims of this study are: to characterize the shell bone microanatomy and osteohistology of different Jurassic, Cretaceous, Cenozoic and extant taxa of Argentina, including members of two main living groups (Pleurodira and Cryptodira); to identify inter- and intra-elemental microanatomical and osteohistological variations in caparace; determine the degree of histological variation between the different Testudinata lineages and to test the taxonomic value of the histological characters; to establish the pattern of shell development in Testudinata; to test whether shell microanatomy is correlated with adaptation to different ecological habitat; to identify possible variations with regard to the shell bone histogenesis among Testudinata and to compare obtained results with previous studies. The main contribution to the paleohistology of shell bone turtle are listed below. First, both intra- and inter-elemetal new histological variation patterns were recorded and previously known patterns were corroborated. New important variation was found in microanatomy and osteohistology between peripheral plates and the rest of the shell plates. Concerning histogenesis, variation on the metaplastic ossification contribution is recorded 4 Tesis Doctoral UNCo J. Marcos Jannello 2018 not only between different shell elements and plate regions, but also between taxa. Metaplastic ossification mainly occurs in external cortex (metaplasic tissue in the internal cortex is present only in few taxa). During ontogeny, metaplastic ossification occurs early or during all ontogeny, but never start at late ontogenetic stages. The different patterns of Testudinata shell development does not exhibit clear phylogenetic signal. A particular growth pattern, where the growth direction is towards internal region (contrary to the most of the Testudinata taxa) is recorded in some species. Two basic mechanisms for external sculpturing formation and maintenance among Testudinata are reported (variation in local rates of bone formation and resorption and new bone tissue deposition). There are not evident relationships between the type and degree of external sculpturing and the mode of its origin and maintenance. Contrary
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